Heteroptera research in Latin America and the Caribbean (Insecta, Hemiptera): status and perspectives in the 21st century.

Latin American and the Caribbean regions (LAC) harbor one of the most biodiverse areas of the world, the Neotropics. True bugs (Hemiptera: Heteroptera) are a diverse lineage of insects, with more than 45,000 species, particularly speciose in the Neotropical region. True bugs are fundamental in the dynamics of natural and modified ecosystems, with several species critical to agriculture and public health. We compiled Heteroptera research in LAC from 1998-2022 using bibliographic databases. Productivity, collaborative networks, and the main topics studied were analyzed. A total of 1,651 Heteroptera studies from LAC were found, with continuous growth being 2021 the most prolific. Four categories (Taxonomy of extant species, Faunistic inventories and new records, Pest species biology, and Community ecology) represent most of the published research. About 60 percent of the records evaluated correspond to five families (Pentatomidae, Reduviidae, Coreidae, Miridae, and Rhyparochromidae). We emphasize the need to keep working on Heteroptera taxonomy because it will allow further advances in other areas such as phylogenetic analyses, biogeography, ecology, and natural history, among others. The results of our analyses characterize the current state of heteropterology in the region, establishing a baseline for future studies and efforts to broaden the knowledge of the group.

Evolutionary mechanisms for camouflage in Cladomorphus phyllinus (Phasmatodea): A reflection on the role of evidence for hypotheses proposition.

We address and discuss some of the many flaws exhibited by Costa et al. (2019) which tried to explain the twig-like camouflage of Cladomorphus phyllinus. Given the lack of both empirical and theoretical underpinnings in Costa et al. (2019), we call into question the validity of their conclusions, in particular, that horizontal gene transfer is a causal mechanism for the camouflage in C. phyllinus.

Taxonomic notes on Ochlerus: revisiting Herrich-Schäffer's species (Hemiptera: Pentatomidae: Discocephalinae: Ochlerini).

The status of the three species of Ochlerus Spinola, 1837, described by Herrich-Schäffer in 1844 is reviewed. Ochlerus coriaceus and O. lutosus are removed from the synonymy of O. cinctus Spinola, 1837, while O. sordidus is transferred to Paralincus sordidus. Lectotypes are designated, and photos for each type and new records for O. coriaceus are provided.

External egg structure of the Pentatomidae (Hemiptera: Heteroptera) and the search for characters with phylogenetic importance.

The chorionic structure of sixteen pentatomid species is described. Morphological patterns in different taxonomic levels are discussed. In addition, egg characters useful in cladistic analyses are listed, and some of those characters are tested for congruence with two cladistic analyses previously conducted within Pentatomidae. Descriptive studies were conducted with Banasa induta, Capivaccius bufo, Catulona pensa, Chinavia armigera, Chinavia aseada, Chinavia brasicola, Chinavia runaspis, Dichelops furcatus, Euschistus heros, Euschistus riograndensis, Euschistus paranticus, Mormidea cornicollis, Podisus distinctus, Podisus nigrispinus, Serdia apicicornis, and Thoreyella maracaja. The eggs were examined and photographed under light and scanning electron microscopy. Based on literature data, a list of 40 egg characters with potential phylogenetic importance has been compiled. Some of these characters were included in the cladistic analyses of the genus Nezara (six characters) and of the Chinavia obstinata group (five characters). Both analyses were performed in TNT with equal weighting of characters. The eggs of most of the Neotropical species studied were barrel-shaped, chorion translucent and spinose, with aero-micropylar processes that were short and clubbed. The patterns of egg morphology could be identified in Carpocorini, Procleticini, and in the genera Banasa, Chinavia, Euschistus, Mormidea, and Podisus. In the cladistic analyses, the inclusion of egg characters did not affect the topology of the trees shown in the original papers. For the analyses, the egg characters were somewhat informative. At present, a total of 286 Pentatomidae species have their egg stage described. 

Paropsocolis, a new pentatomid genus with two new species from Brazil (Heteroptera, Pentatomidae, Ochlerini).

When examining specimens of the Janeirona Distant, 1911 clade of Ochlerini (Discocephalinae) we found males and females of two undescribed species. Though clearly similar to the genera in the Janeirona clade, these two species lack the diagnostic characteristics of any of them. We made a cladistic analysis to verify the placement of the two new species in Discocephalinae. A total of 38 taxa and 60 morphological characters were included in the analysis. Character polarization followed the outgroup comparison method, and the most parsimonious trees were calculated in TNT using an implied weighting approach. The analysis resulted in one most parsimonious tree where the two new species form a monophyletic group within the Ochlerini, sister to Janeirona. We propose Paropsocolis Sampaio & Campos, gen. nov. for the two new species Paropsocolis convergens Sampaio & Campos, sp. nov. and Paropsocolis divergens Sampaio & Campos, sp. nov.. The new genus is recognized by having an inconspicuous labial intercalary segment, metasternum anteriorly carinated, parameres with a basal dish-shaped process and a bifurcated apex, theca with one apical ventral process, endosoma with one dorsal and one ventral processes, laterotergites IX surpassing the mediotergite VIII, and thickening of vaginal intima circular. P. convergens Sampaio & Campos, sp. nov. and P. divergens Sampaio & Campos, sp. nov. are described to Brazil. A distributional map is provided.

Revision of Heteroscelis Latreille, 1829 (Heteroptera: Pentatomidae: Asopinae) with the description of a new species from Colombia.

The Asopinae comprises about 300 species and 63 genera of predatory stinkbugs, of which around 100 species and 23 genera occur in the Neotropical region. The species of the Neotropical genus Heteroscelis Latreille, 1829 are easily recognized by the small size, vivid and often iridescent colors, the elongated head, and the foretibial foliaceous expansions. In this work we provide a taxonomic revision of Heteroscelis. The genus and its four known species are redescribed, and one new species from Colombia, Heteroscelis vandembergi Sampaio & Campos, sp.nov., is proposed. A distribution map, and a key to the species of Heteroscelis are presented.

Morphology of interommatidial sensilla of Discocephalini (Heteroptera: Pentatomidae: Discocephalinae).

Interommatidial sensilla have been explored for their taxonomic value and other aspects in Diptera and Lepidoptera. In Heteroptera, Eurystethus is the only genus where these structures were examined in delimiting subgenera and species. This study investigates the presence, distribution, and morphology of interommatidial sensilla in Discocephalinae species and discusses the sensilla's value for cladistic and taxonomic appraisals. Twenty species of 18 genera were analyzed: 16 Discocephalini genera, one genus in Ochlerini, and one genus in Edessinae. Species' heads were observed from photographs taken with light microscopy (LM) and scanning electron microscopy (SEM). From these images, we identified eleven types of sensilla, classified into chaetica, trichoidea and coeloconica. In LM, only sensilla chaetica with a length of 40-100 µm were visible, present in six species. Under the SEM at least one type of sensillum was visible in 19 of the 20 species analyzed. No difference between the various kinds of sensilla were apparent between males and females or adults and nymphs of a given species. Both subgenera of Eurysthetus have the same type of sensillum (chaeticum type I), which suggests this is not a delimiting feature. We conclude that sensilla chaetica can be a valuable character to delimit groups of genera within Discocephalini.

The eggs and nymphs of predatory stink bugs (Hemiptera: Pentatomidae: Asopinae): what do we know?

The Asopinae are known for their predatory behavior, differing from the phytophagous habits of most pentatomoids, feeding mostly on soft body insects such as larvae of Lepidoptera and Coleoptera. For this reason, asopines have been studied as biological controllers in integrated pest management programs. Notwithstanding their clear relevance, the general knowledge about Asopinae has important gaps, especially regarding immature. Thus, the importance of studying eggs and nymphs of true bugs (Heteroptera) is evident, contributing to understand their classification, biology, and evolution. In this perspective, we conducted a research about immature of predatory stink bugs, highlighting critical features for identification. We present: (1) a literature overview about eggs and nymphs of predatory stink bugs guided by selected categories; (2) images of females laying eggs of ten species and nymphs of thirty-four species, obtained on websites with a citizen science approach; (3) a comparative morphology of immature of six species reared under laboratory conditions, which we examined using light and scanning electron microscopy. We found a remarkable morphological diversity of both eggs and nymphs of Asopinae, revealing key features to establish diagnoses for identification and potential characters to phylogenetics, such as the aero-micropylar processes and chorion scultpturing of the eggs; and the coloration, labium and abdominal plates morphology of nymphs. The results show that little is known about Asopinae immature considering the diversity of the group; however, information obtained by citizen science initiatives, for instance, can improve this knowledge.

The scent of life: Foreword from the Editors.

This special issue of Zootaxa is published in honor of Dr. Jocélia Grazia (Fig. 1) from the Universidade Federal do Rio Grande do Sul, Brazil, one of the world's most respected and influential active taxonomists in Heteroptera. Though she retired from her long-time position as a Full Professor, away from undergraduate classes and administrative duties, Jocélia is still an active researcher and passionate about studying true bugs. An intense conviction she passes to her many students and becomes evident in her publications. If there is one word to define Jocélia Grazia, it is energy. Even the most novice and inexperienced student perceives the enthusiasm and dedication Jocélia faces from the small bureaucratic problems through the most challenging scientific questions.

Revision of emMacropygium/em Spinola, 1837 (Hemiptera: Pentatomidae: Ochlerini) with the revalidation of three species and the description of one new species.

The Ochlerini (Discocephalinae) are endemic in the Neotropical region, and the tribe currently contains 38 genera and 149 species. The monotypic Macropygium Spinola, 1837 is the most widespread ochlerine genus in the Americas, ranging from Mexico to Argentina. Macropygium reticulare (Fabricius, 1803) is one of the few Ochlerini of phytosanitary importance, transmitting Phytomonas spp. in palm crops in South America. Nonetheless its wide distribution, and phytosanitary importance, the taxonomy of Macropygium has been neglected. Here we present a revision of Macropygium, revalidate three species, Macropygium spinolae Stål, 1860, REVISED STATUS (removed from the synonymy with M. reticulare), Macropygium bifidum (Westwood, 1837), NEW COMBINATION (removed from the synonymy with M. reticulare), and Macropygium subsulcatum (Amyot Serville, 1843), REVISED STATUS (removed from the synonymy with M. reticulare). We remove Ochlerus guttipes Walker, 1867 from the synonymy with M. reticulare, and propose O. guttipes a new junior synonym of M. spinolae. Finally, we describe one new species, Macropygium graziae Silva Campos, Sp. Nov.. A key for the species of Macropygium is provided.

Bizarre morphology in extinct Eocene bugs (Heteroptera: Pentatomidae).

Newly discovered fossil bugs (Insecta: Hemiptera: Heteroptera: Pentatomidae) from the Eocene of Messel (Germany) and Green River (North America) exhibit an exaggerated morphology including prominent spiny humeral and anterolateral angles of the pronotum and a spiny lateral abdominal margin. Especially the humeral angles are unique; they consist of expansive, rounded projections with strong spines, which is a rare trait among pentatomids. A hypothesis for the function of this extreme morphology is defence against small vertebrate predators, such as birds or reptiles. The same protuberances also produce a disruptive effect camouflaging the specimen in its environment and provide additional protection. Therefore, the extreme morphology provides primary as well as secondary anti-predator defence. The morphology of Eospinosus peterkulkai gen. et sp. nov. and E. greenriverensis sp. nov. resembles that of Triplatygini, which today occur exclusively in Madagascar, as well as that of Discocephalinae or Cyrtocorinae, which today occur in the Neotropics. Due to a lack of conclusive characters, it cannot be excluded that the fossil species may represent a case of remarkable convergence and are not related to either taxon. Phylogenetic analyses using parsimony as well as Bayesian algorithms confirmed that the new genus is a member of Pentatomidae, but could not solve its phylogenetic relationships within Pentatomidae.

Three new species of emEurystethus/em from Brazil (Hemiptera: Heteroptera: Pentatomidae).

The Heteroptera (true bugs) and its fourth-largest family Pentatomidae (stink bugs) are worldwide distributed groups with higher diversity in the tropics. The pentatomid subfamily Discocephalinae is almost restricted to the Neotropical Region, and it is divided into the tribes Discocephalini and Ochlerini. Eurystethus Mayr, 1864 is placed within Discocephalini and comprises 18 species, 16 out of them classified into two subgenera. Here we describe Eurystethus jo sp. nov., Eurystethus multipunctatus sp. nov., and Eurystethus rufodorsatus sp. nov., all occurring in Brazil, compare them to other species and provide a distribution map.

Taxonomy of widespread Neotropical species of Podisus Herrich-Schäffer (Hemiptera: Pentatomidae: Asopinae): redescription of P. distinctus (Stål, 1860) and revalidation of P. fuscescens (Dallas, 1851).

Some species of Podisus Herrich-Schäffer have a confuse taxonomic history with several modifications and loss of information over time. The genus has been considered a taxonomic dumping ground in Asopinae, and its current species composition is doubtful. After examining the type specimens of Telepta distincta Stål, Arma submarginata Walker, Arma monospila Walker, Arma fuscescens Dallas, Arma caliginosa Walker, Arma turbida Walker and Podisus mellipes Bergroth, we redescribe and revalidate A. fuscescens as Podisus fuscescens rev. stat., previously considered a junior synonym of Podisus sagitta (Fabricius). The taxonomic status, morphology and distribution of P. fuscescens and P. distinctus are updated and illustrated. Arma caliginosa, A. turbida and Podisus mellipes become synonyms of P. fuscescens. Additionally, Arma submarginata Walker is removed from its synonymy with Podisus ventralis (Dallas) and treated as a synonym of P. distinctus. Podisus signoreti Fallou is removed from its synonymy with P. distinctus and treated as species inquirenda.

External morphology of the abdominal glands in Asopinae (Hemiptera: Heteroptera: Pentatomidae).

Heteropterans communicate chemically through thoracic or abdominal glandular complexes. The dorso-abdominal scent glands (DAGs) are externalized by cuticular specialized structures forming the external scent efferent system (SES). Different groups in Heteroptera present other glands, such as the Asopinae (Pentatomidae), whose males can have ventral abdominal pheromonal glands externalized by cuticular modifications, called glandular patches (GPs). Here we describe the SES of DAGs and, for the first time, the GPs of 22 Asopinae genera. The ostioles of SES 1 vary in shape. SES 2 and SES 3 were restricted to scars, indicating that only the SES 1 remains functional in adults. The GPs are found mostly in segments V and VI. We have not found any difference in the morphological pattern between the SES of species bearing GPs and those lacking it, meaning that the acquisition of GPs is likely related to some sexual behavior, thus not as a complex structure replacing any role of the DAGs. The cuticle of the GPs is microsculptured, bearing many pores surrounding the base of sensilla being the pathway for the secretions to reach the outside of the exoskeleton. The sensilla may be adaptations to increase the contact surface, assisting rapid evaporation of the volatile secretions from the pores.

Revision of Tynacantha Dallas with description of two new species (Hemiptera: Heteroptera: Pentatomidae: Asopinae).

Here, Tynacantha Dallas is revised and two new species are described from Brazil: T. cuprea Brugnera Grazia sp. nov. and T. umeridenigrata Brugnera Grazia sp. nov. The type material is analyzed, figures, illustrations, a distributional map and a key to the species are provided. The color variation found in T. marginata is also discussed.

Description of two new genera of Ochlerini (Hemiptera: Pentatomidae: Discocephalinae).

Since the phylogenetic analysis of the Herrichella clade (Ochlerini) new genera were described to fit monophyletic groups inferred and considered to be new taxa. The clade I supported by two synapomorphies of the female genitalia, grouped Alitocoris brunneus Sailer together to two new species. Alloeogyna gen. nov. is described here for this clade including the two new species Alloeogyna ampla sp. nov. and Alloeogyna poecila sp. nov. and a new combination is proposed for Alloeogyna brunnea comb. nov.. Also, Alitocoris maculosus Sailer recovered within the clade E in the Herrichella clade is here redescribed in the monotypic Sailerus gen. nov. resulting in the new combination Sailerus maculosus comb. nov..

Nomenclatural changes in Ochlerini (Hemiptera: Heteroptera: Pentatomidae: Discocephalinae).

Several nomenclatural problems were found while studying the types of Ochlerini in the Natural History Museum, London. These problems are addressed and corrected: Hemingius Distant, 1899 syn. nov. is removed from the synonymy with Moncus Stål, 1867, and considered a new junior synonym of Ochlerus Spinola, 1837; Ochlerus scaber Walker, 1867 is removed from the synonymy with Ochlerus obscurus Dallas, 1851, and it is proposed as a senior synonym of Ochlerus cotylophorus Breddin, 1910 syn. nov. Herrichella Distant, 1911 is a senior synonym of Alitocoris Sailer, 1950 syn. nov., with the resulting new combinations: Herrichella grandis (Garbelotto Campos, 2013) comb. nov., Herrichella lateralis (Garbelotto Campos, 2013) comb. nov., Herrichella ornata (Garbelotto Campos, 2013) comb. nov., and Herrichella schraderi (Sailer, 1950) comb. nov. A key for the species of Herrichella is also given. Eritrachys antennata (Distant, 1911) comb. nov. is a senior synonym of Eritrachys bituberculata Ruckes, 1959 syn. nov. Phereclus punctatus (Dallas, 1851) is transferred to the genus Lincus Stål, 1867 as Lincus punctatus (Dallas, 1851) comb. nov.

Phylogeny and revision of Schraderiellus Rider, 1998 with the description of five new species (Hemiptera, Pentatomidae, Discocephalinae, Ochlerini).

A cladistic analysis of Schraderiellus Rider, 1998 is provided in this study. Were used 60 morphological characters for 34 taxa, including the two known species of Schraderiellus and five new species in the ingroup, two species of Discocephalini and 24 species of Ochlerini in the outgroup, and Euschistus hansi Grazia, 1987 as root. The analysis was made using TNT through heuristic searches under implied weighting of characters. The analysis indicates the monophyly of Schraderiellus with seven valid species. Schraderiellus is redescribed, and five new species are described: S. brevicolis sp. nv., S. falcatus sp. nv., S. luridomaculatus sp. nv., S. luteipedis sp. nv. and S. rufilineatus sp. nv., and a key for the species is presented.

Description of four new species of Ochlerini from South America (Hemiptera: Pentatomidae: Discocephalinae).

The tribe Ochlerini Rolston was proposed in 1981 to house 23 genera of Neotropical pentatomids, most of them containing few species and poorly represented in entomological collections. After a quarter century of accumulated taxonomic knowledge the tribe totals 36 genera and 133 species, but a number of taxa still require revision or remain undescribed. Here we describe and illustrate four new species from South America, Cromata flavida sp. nov., Lincus curvatus sp. nov., Ocellatocoris longirostris sp. nov., and Xynocoris ecuadorensis sp. nov. The specimens of L. curvatus and X. ecuadorensis were collected on oil palms (OxG hybrids, Elaeis oleifera x Elaeis guineensis) affected by the lethal wilt (marchitez sorpresiva) disease.

Unusual looking pentatomids: reassessing the taxonomy of Braunus Distant and Lojus McDonald (Hemiptera: Heteroptera: Pentatomidae).

The Neotropical genus Braunus Distant is revised. Based on general morphology, Braunus is transferred to the Pentatominae and Lojus is proposed as a junior synonym of Braunus. The male and female genitalia of Braunus sciocorinus (Walker) are described for the first time. Three new species from the Peruvian and Bolivian Andes and from Colombia are described, Braunus machadoi Barão & Garbelotto sp. nov. (Holotype male deposited in AMNH: Peru, Cuzco, 01.XII.2011, C. Weirauch leg.), Braunus gibbus Garbelotto & Barão sp. nov. (Holotype female deposited in AMNH: Bolivia, Santa Cruz, Prov. Caballero, 15-21.X.2001, S. Spector & J. Ledezma leg.), and Braunus prionotus Barão & Garbelotto sp. nov. (Holotype female deposited in MNHN: Colombia, VII.1974, Steinheil leg.), respectively. Also, we describe the internal male and female genitalia of Braunus ocellatus (Thomas) comb. nov. A key to species and comments on subfamilial and tribal placements and on wing polymorphism are provided.